Life history and spatial distribution of Oriental beetle (Coleoptera: Scarabaeidae) in golf courses in Korea

Larval and adult activity of the oriental beetle Exomala orientalis (Waterhouse), a pest of turfgrass in Korea, was investigated at four golf clubs in Pusan, Korea, from 1995 to 1999. Adult emergence was first observed on the greens in late May with peak activity occurring 2 wk later. During the day, E. orientalis adults were most active between 1800 and 2200 hours. First instars were found mostly in early July, second instars mostly in late July, and third instars from August to April. The density of larvae in fixed plots decreased steadily from the time of egg laying to pupation: 667/m3 on 26 July, 267/m3 on 29 August, and 122/m3 on 2 October 1997. All the observed E. orientalis completed one generation per year. Adult females were observed feeding on flowers of a late-blooming variety of Japanese chestnut (Castanea crenata Sieb & Zucc). E. orientalis larval densities were higher in greens with Japanese chestnut nearby, and where magpie, Pica pica sericea (Gould), feeding was observed. More E. orientalis adults emerged from the right, left, and back of greens than from the front or middle. The intensity of emergence was inversely proportional to the amount of golfer traffic on various parts of the green. Counting emergence holes may be a way that golf course superintendents can predict which greens and tees are most likely to be damaged from E. orientalis larvae without destructive sampling.     

Life history and population dynamics of Ectinohoplia rufipes (Coleoptera: Scarabaeidae) on Korean golf courses

Life history and temporal distribution of Ectinohoplia rufipes larvae (Coleoptera: Scarabaeidae; Melolonthinae) were studied on multiple courses at two Korean golf clubs in Gapyeong and Anseong, Gyeonggi Provinces. Larval density of E. rufipes was higher in roughs than fairways and higher on older courses than on newly constructed ones at each club. Larval density of E. rufipes in fall reached three to four times compared with the previous year in five or six year old courses in the first opened courses, but that took two or three years to reach three to four times in the newly constructed courses in the same golf club. Monthly sampling showed that larvae were distributed 3.0 to 5.8 cm beneath the soil surface; mean larval density ranged from 0.2 to 1.2/m² at each club. Most larvae overwintered as third instars that pupated in early May. Adults were first observed in late May at each course. Adults were first seen in emergence cages installed over a grub-infested portion of rough in early June. Eggs were deposited from late June to early July. Eggs hatched in mid to late July and larvae developed to become third instars by early October. Implications of these findings for managing E. rufipes on Korean golf courses are discussed.     

Male and female Popillia quadriguttata (Fabricius) and Protaetia brevitarsis (Lewis) (Coleoptera: Scarabaeidae) response to Japanese beetle floral and pheromone lures

Popillia quadriguttata (Fabricius), and Protaetia brevitarsis (Lewis) adults were captured with Japanese beetle, Popillia japonica Newman, sex attractant and floral lures at Changchun, China during July–August 2012. The floral lure (phenethyl propionate:eugenol:geraniol, 3:7:3) was attractive to male and female P. quadriguttata (AV: 1.2 ± 0.9; 1.1 ± 0.3; total: 2.3 ± 0.8), and was similar in attraction to the combination of the sex attractant (SA) [(R, Z)-5-(1-decenyl) dihydro-2(3H)-furanone] plus the floral lure for male (1.60 ± 0.2), female (1.30 ± 1.1) and total captures (2.9 ± 3.0). However, the SA alone captured only males in much higher numbers than when combined with the floral lure (10.0 ± 6.4). In a separate earlier test, the greatest number of P. quadriguttata males (12.5 ± 3.0), female (12.2 ± 1.5) and total captures (24.7 ± 2.5) was in yellow, laboratory-made, bottle traps. The floral lure also attracted female Pro. brevitarsis (10.0 ± 3.4), while the SA attracted only few male beetles (1.0 ± 0.2). The combination SA + floral lure captured similar females (11.0 ± 2.0) and total (14.2 ± 2.2) Pro. brevitarsis as the floral lure alone. Two butterflies, Colias erate poliographus (Motschulsky) and Pieris rapae (Linnaeus), were also attracted to the floral lure. These studies indicate a potential for replacing pesticides by using the Japanese beetle lures for monitoring and control of several insects in China, and that they would be useful in monitoring and eradication of two potential scarab pests, P. quadriguttata and Pro. brevitaris, in the United States and Europe.     

Incidence of turf-damaging white grubs (Coleoptera: Scarabaeidae) and associated pathogens and parasitoids on Kentucky golf courses

Root-feeding grubs (Coleoptera: Scarabaeidae) were sampled from damaged areas of 61 irrigated roughs on 32 Kentucky golf courses to determine species composition and natural enemy incidence, the first such survey in the United States' transitional turfgrass climatic zone. Masked chafers (Cyclocephala lurida Bland and C. borealis Arrow) and Japanese beetle (Popillia japonica Newman) accounted for ≈73 and 26% of grubs found in an autumn survey, with Cyclocephala spp. predominating at most sites, although mixed infestations were common. Only a few Phyllophaga spp., and no exotic species other than P. japonica were found. Cyclocephala spp. also predominated in seasonal and statewide surveys regardless of whether a course had cool- or warm-season grass fairways. Pathogenic bacteria, Paenibacillus and Serratia spp., and the autumn-active parasitoid Tiphia pygidialis Allen were the main enemies associated with Cyclocephala spp. Predominant enemies of P. japonica were Paenibacillus, Serratia, and Metarhizium spp. in autumn, and eugregarines, Stictospora sp. (probably S. villani Hays and Clopton) and Tiphia vernalis Rohwer in spring. Entomopathogenic nematodes and the microsporidian Ovavesicula popilliae Andreadis & Hanula were nearly absent in our samples. No predictive relationships were found between soil parameters and proportionate abundance of Cyclocephala or P. japonica, or with natural enemy incidence at particular sites. Although incidence of individual enemies was generally low (<20%; often <5%) in these point-in-time surveys, collectively and over their hosts' prolonged development they may take a significant toll on grub populations.     

Dispersal of Steinernema glaseri (Nematoda: Steinernematidae) in adult Japanese beetles, Popillia japonica (Coleoptera: Scarabaeidae)

Japanese beetle adults, Popillia japonica, can become infected with and disperse the entomopathogenic nematode, Steinernema glaseri, under laboratory and field conditions. After a 24-h exposure to 10 000 infective juveniles/20 adult beetles, 45% of the beetles died within 4 days post-treatment, but only 59% of these were infected with the nematode. Corresponding control mortality was 6.5%. An average of 238 infective juveniles were produced/beetle. Beetles exposed to 4000 and 10 000 infectives/10 adults carried with them an average of 17 and 59 infectives/adult on external body surfaces respectively. When beetles that had been exposed to 4000 infectives/20 adults were transferred to, and held in, cages containing soil for 2 weeks, up to 89% of the adults died, as did 74% of the P. japonica larvae that were subsequently placed in the cages. When adults that had been exposed to 50 000 infectives/250 beetles in moist sand for 16 h were released into screened cages in the field at soil temperatures of over 25 C, the soil beneath 83% of the cages tested positive for the nematode, using Galleria mellonella larvae as bait, 2 weeks after releasing the beetles. No nematodes were detected in control plots. The potential of infected adult P. japonica for dispersing S. glaseri by flight was investigated by exposing adults to 50 000 infectives/250 beetles, marking and releasing them in the field and recapturing them in lure-baited Japanese beetle traps. Less than 1% of the treated beetles were recaptured, but 33% of these had one or more nematodes in their hemocoels. Accordingly, this approach does not appear to be feasible for large-scale augmentation and dispersal of the nematode using currently developed methods of infection. If improvements in mass-inoculation methods can be made that enable a rapid high percentage of infection while still permitting flight, this concept could be employed to establish new foci of infection or for the introduction of other species of nematodes.     

Survey of pathogens and parasitoids of Popillia japonica (Coleoptera: Scarabaeidae) in northwest Arkansas

The impact of pathogens and parasitoids on the recently established population of Popillia japonica Newman in northwest Arkansas has been unknown. In this study, we quantified the prevalence of natural enemies: Stictospora villani Hays, Ovavesicula popilliae Andreadis and Hanula, Paenibacillus spp. (Dingman), nematodes and parasitic Diptera and Hymenoptera in third instar and adult populations in 2010 and 2011. S. villani was found in 38.4% and 35.5% of larvae in 2010 and 2011, respectively. S. villani was not found in adult beetles. Paenibacillus bacteria were not found in either larvae or adults in either year. In 2010, the microsporidian O. popilliae was not found in larvae but was present in 0.2% of adults. In 2011, 2.6% of larvae were infected with O. popilliae, but the microsporidian was not found in adults. A previously unknown Adelina sp. was found infecting 0.4% of adult beetles in 2010 and 1.3% of larvae in 2011. Nematode infections were found in 1.8% of larvae and 0.1% of adults in 2010 and not found in either life stage in 2011. No parasitic Hymenoptera or Diptera were found in either year. Apparently, pathogens and parasitoids currently provide little control of the Japanese beetle population within northwest Arkansas.     

How the use of chlorothalonil on golf courses impacts Paenibacillus sp., a pathogen of Ataenius spretulus (Coleoptera: Scarabaeidae)

One of the most frequently used fungicides on golf courses, chlorothalonil, was applied to field plots at four different golf courses to determine how Ataenius spretulus Haldeman, Aphodius granarius L., carabids, staphylinids, formicids and Paenibacillus sp. were affected. Chlorothalonil had no influence on the incidence of Paenibacillus sp. infection of A. spretulus larvae in golf course fairways. The incidence of Paenibacillus sp. infection of A. spretulus larvae was much greater in the rough (47.4%) compared with the fairway (26.4%) at Cattails Golf Club despite a more dense population (fourfold) of larvae in the fairway. Chlorothalonil treatment of turf plots had no influence on the number of staphylinids, carabids, formicids, or A. spretulus adults caught in pitfall traps. It also did not affect the density of A. spretulus larvae. Data from this study suggest that the widespread use of chlorothalonil on fairways is not likely to cause outbreaks of A. spretulus on golf courses.     

Timing of onset of evening activity of adult Chinese rose beetles (Coleoptera: Scarabaeidae)

Adult Chinese rose beetles, Adoretus sinicus (Burmeister) (Coleoptera: Scarabaeidae: Adoretini), present in China, Taiwan, Indonesia, Cambodia, Laos, Singapore, Thailand, Vietnam, the Marianas Islands, the Caroline Islands, and the Hawaiian Islands, are nighttime defoliators that feed on a wide variety of plant species. It has recently been demonstrated that illumination of plants at dusk has the potential to discourage feeding by adult Chinese rose beetles on the illuminated plants. To effectively use lighting to minimize defoliation of host plants, it is critical to know the timing of the initial host plant colonization by the beetles to ensure that illumination is initiated before host plant colonization begins. Adult Chinese rose beetles were observed to colonize host plants at dusk, with initiation of beetle colonization averaging more than 21 min after sunset, with the earliest observed beetle colonization occurring 11 min after sunset. These times corresponded to an average light level of 7.0 lux at the first colonization and the earliest first colonization occurring at 26.9 lux. Based on these results, use of lighting to minimize defoliation of host plants should be initiated at about sunset in order to discourage colonization (and associated defoliation) by adult Chinese rose beetles.     

Ultrastructure of pheromone-detecting sensillum placodeum of the Japanese beetle, Popillia japonica Newmann (Coleoptera: Scarabaeidae)

The pheromone-detecting sensilla placodea are significantly more numerous than other sensory structures in the antennae of the Japanese beetle, Popillia japonica (Coleoptera: Scarabaeidae). Their abundance in males is nearly twice of that in females, showing a clear sexual dimorphism. Externally, they have a tortoise shell-like round cuticular plate containing a few polygonal plates separated by narrow ridges. Internally, they house two long dendrites that branch and terminate near fine cuticular pores. They have a system of two bipolar neurons accompanied by three enveloping cells, resembling sensilla trichodea in moths. The conspicuous difference with the latter is that the sensillum-lymph cavity near the outer cuticle is funnel-shaped, into which the tormogen cell projects numerous microvilli whose tips approach the terminal branches of the dendrites.     

Crepuscular activity of adult Sericesthis geminata (Coleoptera: Scarabaeidae): Influence of circadian rhythmicity and light intensity

Subterranean Sericesthis geminata (Boisduval) beetles emerge from the soil daily during the flight season. Emergence and flight are initiated within 15 min after sunset. The beetles are active above the soil surface as the intensity of illumination decreases from 240 lux to 0.7 lux.

If young adult beetles still in their pupal cells in the soil are placed under constant, low‐intensity illumination, they first emerge at any time of day, showing no cyclical diurnal activity pattern. When such beetles are exposed to diurnal fluctuations in illuminance, the first emergence and flight occur at dusk. These beetles subsequently show a diurnal rhythm in their activity, even when exposed to constant environmental conditions. This endogenous activity cycle has a period of about 22 h, and is synchronised with diurnal fluctuations in light intensity. It is reset in response to a change in the photoperiod.

The data suggest that, after initial emergence from the soil, daily crepuscular activity is initiated by an endogenously controlled activity rhythm which causes the beetles to burrow to the soil surface shortly before dusk. At the soil surface they are exposed to fluctuations in the duration and intensity of illumination which may reset the endogenous rhythm, affecting the time of subsequent emergence.     

Entomopathogenic nematodes as potential biological control agents of Popillia japonica (Coleoptera,Scarabaeidae) in Piedmont Region (Italy)

In 2014, the Japanese beetle Popillia japonica (Coleoptera: Scarabaeidae) was first spotted in northern Italy in the Nature Park of the Ticino Valley, its first detection in continental Europe. This polyphagous invasive species has the potential to cause serious losses to horticulture and agriculture. Particularly for its management in a Nature Park, environmentally friendly strategies are necessary. To develop baseline data for a biological control approach to the Italian outbreak of P. japonica, we conducted laboratory and field experiments testing several indigenous and commercial strains of the entomopathogenic nematode (EPN) species Heterorhabditis bacteriophora and Steinernema carpocapsae against P. japonica larvae. In the laboratory, strains of H. bacteriophora caused greater mortality (ranging from 57% to 100%) than those of S. carpocapsae (3% to 77%). In micro‐plot field tests carried out at three different times, the most virulent indigenous EPN strain, H. bacteriophora ItH‐LU1, showed again the best results ranging from 44% to 93% against young larvae. Finally, in a large‐plot field trial, the commercial H. bacteriophora product provided 46% larval mortality. This study shows that H. bacteriophora strains have good potential as biological control agents of larvae of the invasive P. japonica in northern Italy.     

Developmental stages and gut microenvironments influence gut microbiota dynamics in the invasive beetle Popillia japonica Newman (Coleoptera: Scarabaeidae)

Popillia japonica Newman (Coleoptera: Scarabaeidae) is a highly polyphagous invasive beetle originating from Japan. This insect is highly resilient and able to rapidly adapt to new vegetation. Insect-associated microorganisms can play important roles in insect physiology, helping their hosts to adapt to changing conditions and potentially contributing to an insect's invasive potential. Such symbiotic bacteria can be part of a core microbiota that is stably transmitted throughout the host's life cycle or selectively recruited from the environment at each developmental stage. The aim of this study was to investigate the origin, stability and turnover of the bacterial communities associated with an invasive population of P. japonica from Italy. Our results demonstrate that soil microbes represent an important source of gut bacteria for P. japonica larvae, but as the insect develops, its gut microbiota richness and diversity decreased substantially, paralleled by changes in community composition. Notably, only 16.75% of the soil bacteria present in larvae are maintained until the adult stage. We further identified the micro-environments of different gut sections as an important factor shaping microbiota composition in this species, likely due to differences in pH, oxygen availability and redox potential. In addition, P. japonica also harboured a stable bacterial community across all developmental stages, consisting of taxa well known for the degradation of plant material, namely the families Ruminococcacae, Christensenellaceae and Lachnospiraceae. Interestingly, the family Christensenallaceae had so far been observed exclusively in humans. However, the Christensenellaceae operational taxonomic units found in P. japonica belong to different taxonomic clades within this family.     

Structure and function of the mandibles of adult dung beetles (Coleoptera : Scarabaeidae)

The mouthparts of adult dung beetles (Coleoptera : Scarabaeidae) are adapted for manipulation of soft, pasty food—usually the excrement of vertebrates. Nutrients are derived primarily from micro-organisms contained in the food and these must be broken up before ingestion. The mandibles, particularly the molar lobes, are designed to finely grind these particulates; the molae function as a mortar-pestle system, which mills the organic “grist” contained in the food by a combination of squeezing and grinding actions. The ability of the molae to finely grind food particles resides primarily in the structure of the molar surface, which consists of a series of ridges bearing rows of submicroscopic scrapers (“tritors”). The incisor lobes of the mandibles scrape food from the surfaces of the galeae and lacinae, which bring food into the preoral cavity. The structure of the mouthparts of Canthon pilularius (L.), which is described in detail, is basically the same as that of 10 other species examined. Those of Aphodius and Geotrupes are similar in some respects to those of scarabaeines but are obviously less well specialized for microphagy.     

Two Newly Recorded Coprophagid Species from Korea (Coleoptera, Scarabaeidae and Aphodiidae)

A species of Scarabaeidae, Panelus parvulus Waterhouse and a species of Aphodiidae, Aphodius fossor (Linné) are newly recorded from Korea. The diagnosis and photographs of them are provided.     

An Attractant Trap for Autodissemination of Entomopathogenic Fungi into Populations of the Japanese Beetle Popillia japonica (Coleoptera: Scarabaeidae)

Autodissemination may be effective against the Japanese beetle, Popillia japonica Newman, in situations where habitats of its larvae are inaccessible. Trapping systems with attractants for both male and female Japanese beetles are commercially available. We fabricated an inoculation chamber which fits between the top of a standard Trece Catch CanTM Japanese beetle Trap and its holding canister. Beetles which are attracted to the trap fall through a hole in the inoculation chamber and land on a mesh screen. A partial funnel and canister attachment from a metal Ellisco Japanese Beetle Trap was secured beneath a hole in the floor at the opposite end of the chamber. A 10-cm section in the middle of the box, between the entrance hole in the roof and the exit hole in the floor, allows space for a dish containing the inoculum to be placed into the chamber through a door in the side of the unit. The trap has been tested with Metarhizium anisopliae (Metschnikoff) Sorokin as the pathogen. Beetles emerging from the device in the field were captured and returned to the laboratory where the presence of conidia and mortality to adult beetles from the fungus were confirmed.     

Rearing Costelytra zealandica (Coleoptera: Scarabaeidae)

Interaction between larvae can be a cause of mortality when scarabaeid larvae are concentrated in a confined volume of soil. Larvae of Costelytra zealandica (White), the New Zealand grass grub, were held for 30 days at population densities between 1 and 50 larvae per 200 ml of medium, and the effect of crowding on their survival and weight change was assessed. Larval density had no effect on survival in soil plus chopped sheep dung (3 :1), with up to 10 2nd‐instar larvae or 5 3rd‐instar larvae per 200 ml of medium. Larval survival and weight gain were increased by adding germinating ryegrass seed (Grasslands Ruanui) to the medium base, but the effect of density did not diminish. Weight gain of 3rd‐instar larvae decreased as their density increased. When the amount of germinating ryegrass seed in the medium was varied, there was no significant difference between the survival of larvae held in treatments with 5, 25, and 50 g of seed per litre of medium. Survival was significantly lower with 0 and 100 g of seed per litre of medium. It is concluded that, in rearing conditions, the final density of fully grown 3rd‐instar larvae should be a maximum of about 20 per litre of medium to prevent mortality caused by larval interaction—provided that adequate food is available.     

Morphology and Distribution of Sensilla on the Antennae of Hylamorpha elegans Burmeister (Coleoptera: Scarabaeidae)

Considering that sensilla constitute important functional elements of sensory systems in insects, the aim of this study was to determine the type and distribution of sensilla in the antennae of Hylamorpha elegans Burmeister examined by scanning electron microscopy. Hylamorpha elegans antennae are lamellate and consist of the scape, pedicel, and flagellum. The antennal club of this beetle consists of three terminal plates: proximal, middle, and distal lamellae. Four types of sensilla were observed in the lamellae from both sexes: sensilla trichoidea, chaetica, coeloconica, and placodea. Antennal length was larger in males than in females, and significant sexual variation in the number of sensilla placodea and sensilla coeloconica was observed.     

Effect of insecticides on Tiphia vernalis (Hymenoptera: Tiphiidae) oviposition and survival of progeny to cocoon stage when parasitizing Popillia japonica (Coleoptera: Scarabaeidae) larvae

The effect of insecticides on oviposition of Tiphia vernalis Rohwer and subsequent survival of parasitoid progeny to the cocoon stage was determined in the laboratory by using larval Japanese beetle, Popillia japonica Newman, as the host. Insecticides tested were imidacloprid, thiamethoxam, halofenozide, chlorpyrifos, and carbaryl at labeled rates. Female T. vernalis were allowed 2 d to parasitize P. japonica larvae after the parasitoids had received a 4-d exposure to insecticide-treated soil. Another group of female T. vernalis were allowed 2 d to parasitize P. japonica larvae that had been exposed to insecticide-treated soil for 3-4 d. Percentage of parasitism of P. japonica larvae in these trials after exposure of adult parasitoids to carbaryl, chlorpyrifos, halofenozide, or imidacloprid-treated soil (23.3-50.0%) or adult parasitoids to chlorpyrifos, halofenozide, or imidacloprid-treated grubs (33.0-56.7%) was not negatively affected relative to the control treatment (21.7-54.2%). A third group of adult T. vernalis and P. japonica larvae were simultaneously exposed to chlorpyrifos or carbaryl treatments. Percentage parasitism in these trials was lower for T. vernalis adults exposed to the chlorpyrifos and carbaryl (15.0-25.0%) relative to the control (57.5-62.5%) with the exception of one trial with carbaryl (40.0%). However, exposure of the parasitoid and P. japonica to chlorpyrifos 0.5X, carbaryl 0.5X, imidacloprid, halofenozide, or thiamethoxam in several trials resulted in parasitism that was equivalent or greater than (45.0-80.0%) the untreated control (57.5-62.5%). Japanese beetle larval mortality in these trials was greater in the insecticide and parasitoid combination (97.5-100.0%) than with insecticides alone (45.0-100.0%). Percentage of survival of T. vernalis progeny to the cocoon stage was not negatively affected by a 4-d adult parasitoid exposure to carbaryl and chlorpyrifos treated soil (11.7-16.7% versus 18.3% control) or a 2-d exposure to P. japonica-treated larvae (16.7-18.3% versus 28.3% control). However, simultaneous exposure of T. vernalis progeny and P. japonica larvae to chlorpyrifos- and carbaryl-treated soil resulted in no parasitoids surviving to the cocoon stage. Between neonicotinoids, thiamethoxam had more adverse impact on percentage parasitism (52.5%) and survival to the cocoon stage (10.0%) than imidacloprid (80.0 and 32.5%, respectively). Results of this study indicate soil incorporation of imidacloprid and halofenozide had minimal effect on the number of P. japonica larvae parasitized by T. vernalis or survival of T. vernalis progeny to the cocoon stage; therefore, they are more suitable for use with T. vernalis. In contrast, chlorpyrifos, carbaryl, and thiamethoxam lowered the number of T. vernalis progeny surviving to the cocoon stage, and carbaryl and chlorpyrifos reduced the number of P. japonica larvae parasitized. The soil incorporation of insecticides is discussed as one explanation for the minimal effects of some insecticides on T. vernalis.     

Ecological methods for adult populations of Oryctes rhinoceros (Coleoptera, Scarabaeidae)

ABSTRACT. 1. For comparing assessment methods, O.rhinoceros (L.) populations were monitored in five 30–50 ha plots in southern Luzon, Philippines. No consistent correlations were found between number of beetles caught in traps, amount of palm damage and number of breeding sites. Apparently, plots were too small to account for fast dispersal of beetles.
2. Coconut cap traps baited with ethylchrysanthemumate attracted O.rhinoceros adults searching for breeding sites. Several factors were identified influencing catches but trapping was found to be unsuitable for accurately assessing pest populations.
3. A new method which gives estimates on the monthly number of O.rhinoceros attacks on coconut palms, was tested in the Maldives. Palms are climbed about once a year and the sequence of fronds, the number and the position of beetle cuts are recorded.
4. Reduction with age of the length of three teeth on the fore tibia of O. rhinoceros adults was studied in the Philippines. The data was insufficient to obtain a clear correlation between age and length of teeth.
5. The observations indicated that in the Philippines after leaving their site of pupation, O.rhinoceros adults spend about 5 weeks feeding on coconut palms. This is followed by a period of about 7 weeks in breeding sites and, on occasion, additional visits to palms. With these estimates it was possible to relate palm damage records with numbers of feeding adults.     

Morphological variation in adult Xylotrupes gideon australicus (Scarabaeidae,Coleoptera, Insecta)

Variation in horn and body size of both sexes of Xylotrupes gideon australicus is normally distributed with no evidence of dimorphism in either sex. This contrasts with that of X. gideon gideon, in which males are dimorphic for horn length, but horn size varies within morphs. Males of X. gideon australicus are generally larger than females, but the distributions of all characters, except tarsal length, overlap considerably. Males have longer legs, presumably to facilitate mating.